Isognathotermes fungifaber (Sjöstedt, 1896)

Figs 1, 3–4, 6, 26–31, 33, 42–44, distribution map: Fig. 45; Tables 5–6

Eutermes fungifaber Sjöstedt, 1896: 297 (alate imago only).

Mirotermes (Cubitermes) schmidti Emerson, 1928: 520–521, fig. 62.

Mirotermes (Cubitermes) banksi Emerson, 1928: 522–523, fig. 63.

Mirotermes (Cubitermes) comstocki Emerson, 1928: 525, fig. 64.

Eutermes fungifaber – Sjöstedt 1900: 143–149, pl. IV (all castes).

Mirotermes (Cubitermes) fungifaber — Sjöstedt 1913: 369, pl X, 3. — Emerson 1928: 517–519, fig. 60.

Cubitermes fungifaber – Silvestri 1914: 91. — Sjöstedt 1926: 218–225. — Snyder 1949: 159. — Bouillon & Vincke 1971: 269. — Krishna et al. 2013: 1921–1922. — Josens & Deligne 2019: 60.

Cubitermes banksi – Ruelle 1992: 501. — Krishna et al. 2013: 1913.

Cubitermes comstocki – Ruelle 1992: 501. — Krishna et al. 2013: 1918.

Cubitermes schmidti – Ruelle 1992: 501. — Krishna et al. 2013: 1935.

Cubitermes sp. affinis subarquatus “spA” – Roy et al. 2006: 4–5.

Cubitermes sp. affinis subarquatus “spD” – Roy et al. 2006: 4–5.

Isognathotermes fungifaber – Hellemans et al. 2021: 233.

Diagnosis

The worker has a fungifaber EVA (Fig. 4) but in 2% of the samples examined, it has an EVA intermediate between the fungifaber and finitimus patterns. The worker itself and its enteric valve are among the smallest in the genus Isognathotermes (Fig. 31).

The soldier is among the smallest in the genus Isognathotermes; (SHdL = 2.37–3.11 mm, Fig. 28); its EVA generally looks like a muneris EVA and is among the smallest in the genus Isognathotermes EVAs. In some samples, however, the soldier is as large as the smallest soldiers of I. severus . The soldier has, along with I. planifrons, the most evenly curved mandibles from base to tip: ln(SMlpR) - ln(SMldR) = 0.85–1.88.

The imago is, on average and with that of I. phalloides sp. nov., the smallest in the genus Isognathotermes; (IHdW = 1.40–1.61 mm, Fig. 26).

This species is also defined by its ecology and chorology: it has always been found in evergreen forests from RCI to Gabon at less than 300 km from the Atlantic coast (Fig. 45 but see later, ‘Supplementary material of dubious I. fungifaber ’).

Etymology

The name fungifaber from the Latin fungus (mushroom) and faber (artisan) doubtless refers to the building of mushroom-shaped epigeous nests.

Material examined

One hundred twenty-four samples from 44 locations (plus six samples of “dubious I. fungifaber ” from five locations). Of the 64 old museum samples examined, 49 were initially labelled as C. fungifaber, one as C. fungifaber var. elongata, four as C. banksi, one as C. comstocki, three as C. gaigei, three as C. kemneri, two as C. modestior, and one as C. schmidti . Of the six samples of “dubious I. fungifaber ”, four were initially labelled as C. fungifaber and two not identified.

This species also includes C. aff. subarquatus “spA” and “spD” (in Roy et al. 2006).

Syntypes of Eutermes fungifaber Sjöstedt, 1896

CAMEROON • soldier, worker, ♀ (alate), ♀ (queen); Mbongé; 4°32.18′ N, 9°6.67′ E; Oct. 1891; Y. Sjöstedt leg.; study code: DJ 0271; initially Eutermes fungifaber; NHMM • soldier, worker, ♀ (queen); Mbongé; 4°32.18′ N, 9°6.67′ E; 27 Oct. 1891; Y. Sjöstedt leg.; study code: DJ 0272; initially Eutermes fungifaber; AMNH • worker, ♀ (alate); Mbongé (?); 4°32.18′ N, 9°6.67′ E; 1891; Y. Sjöstedt leg.; study code: DJ 0965; initially Eutermes fungifaber ex typis; RBINS .

Krishna et al. (2013: 1921) mention that other syntype samples are deposited in NHRM, not examined.

Paratypes of Mirotermes (Cubitermes) banksi Emerson, 1928

CAMEROON • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0287; initially Mirotermes (C.) banksi → C. fungifaber; AMNH • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0097; initially C. banksi in RMCA; BE RMCA INS.Iso.059210.

Krishna et al. (2013: 1913) mention that the holotype and paratypes are deposited in UMMZ, not examined.

Paratypes of Mirotermes (Cubitermes) comstocki Emerson, 1928

CAMEROON • soldier, worker, ♀ (queen); Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0273; initially Mirotermes (C.) comstocki → C. fungifaber; AMNH • soldier, worker, ♀ (alate); Bipindi?; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0278; initially C. comstocki → C. fungifaber; AMNH • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0121; initially C. comstocki in RMCA; BE RMCA INS.Iso.059212.

Krishna et al. (2013: 1918) mention that the holotype (soldier) and paratypes are deposited in UMMZ, not examined, and that other paratypes are deposited in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, not examined.

Paratypes of Mirotermes (Cubitermes) schmidti Emerson, 1928

CAMEROON • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0275; initially C. schmidti → C. fungifaber in AMNH • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0124; initially Mirotermes (C.) schmidti in RMCA; BE RMCA INS. Iso.059213

Other material examined

CAMEROON • soldier; Abonando [= Ossidinge]; 5°54′ N, 9°8′ E; no date; G. Mansfeld leg.; study code: DJ N123; initially C. fungifaber; AMNH • soldier, worker, ♀ (alate); Mungo; 5°12′ N, 9°31′ E; 1903; Rohde leg.; study code: DJ 0269; initially C. fungifaber var. elongata; AMNH • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0963; initially Eutermes fungifaber; RBINS • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 1920; G. Zenker leg.; study code: DJ 0964; initially Eutermes fungifaber; RBINS • soldier; Réserve du Nyong; 3°50′ N, 10°50′ E; 6 Dec. 1949; J. Birkett-Smith and J. Dahl leg.; study code: DJ U036; initially C. fungifaber; NHMUK 13671854 • soldier; Edea; 3°29′ N, 9°50′ E; Jul.–Sep. 1973; N.M. Collins leg.; study code: DJ U033; initially C. fungifaber; NHMUK 13671851 • soldier; Edea; 3°29′ N, 9°50′ E; Jul.–Sep. 1973; N.M. Collins leg.; study code: DJ U039; initially C. fungifaber; NHMUK 13671847 • soldier; Edea; 3°29′ N, 9°50′ E; Jul.– Sep. 1973; N.M. Collins leg.; study code: DJ U040; initially C. fungifaber; NHMUK 13671850 • soldier; Edea; 3°29′ N, 9°50′ E; Jul.–Sep. 1973; N.M. Collins leg.; study code: DJ U041; initially C. fungifaber; NHMUK 13671848 • soldier; Edea; 3°29′ N, 9°50′ E; Jul.–Sep. 1973; N.M. Collins leg.; study code: DJ U042; initially C. fungifaber; NHMUK 13671846 • soldier; Mbalmayo; 3°30′ N, 11°31′ E; 1 Dec. 1962; G. Becker leg.; study code: DJ U065; initially C. fungifaber; NHMUK 13671852 • soldier, worker; Mamfe–Ikom road; 5°47′ N, 8°58′ E; 18 Sep. 1966; W. Wilkinson leg.; study code: DJ 0718; initially C. gaigei; NHMUK 13671886 • worker, ♂ (king); Edea; 3°29′ N, 9°50′ E; Jul.–Sep. 1973; N.M. Collins leg.; study code: DJ 0649; initially C. fungifaber; NHMUK 13671846 • soldier, worker, ♀ (alate); Ebodjié; 2°38′ N, 9°53′ E; Nov. 1991; A. Dejean leg.; study code: DJ 0096; initially C. banksi in RMCA; BE RMCA INS.Iso.059211 • soldier; Akok; 3°53′ N, 11°57′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U049; initially C. fungifaber; NHMUK 13671877 • soldier; Akok; 3°53′ N, 11°57′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U050; initially C. fungifaber; NHMUK 13671876 • soldier; Akok; 3°53′ N, 11°57′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U052; initially C. fungifaber; NHMUK 13671879 • soldier; Akok; 3°53′ N, 11°57′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U053; initially C. fungifaber; NHMUK 13671880 • soldier, ♀ (alate); Ebogo; 3°23′ N, 11°28′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U045; initially C. fungifaber; NHMUK 13671874 • soldier; Ebogo; 3°23.5′ N, 11°28′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U068; initially C. kemneri; NHMUK 13672024 • soldier; Ebogo; 3°23.5′ N, 11°28′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U069; initially C. kemneri; NHMUK 13672025 • soldier; Ebogo; 3°31′ N, 11°30′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U092; initially C. banksi; NHMUK 13671831 • soldier, ♂ (king), ♀ (queen); Eboufek; 3°31′ N, 11°30′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U051; initially C. fungifaber; NHMUK 13671875 • soldier, worker, ♂ (king); Eboufek; 3°30′ N, 11°53′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0651; initially C. kemneri; NHMUK 13672023 • soldier, worker, ♂ (alate); Eboufek; 3°30′ N, 11°53′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0707; initially C. banksi; NHMUK 13671828 • soldier, ♂ (king); Ekombitié; 3°28′ N, 11°31′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U048; initially C. fungifaber; NHMUK 13671872 • soldier; Mbalmayo; 3°27′ N, 11°29′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U047; initially C. fungifaber; NHMUK 13671871 • soldier; Mbalmayo; 3°27′ N, 11°29′ E; Feb. 1996; P. Eggleton et al. leg.; study code: DJ U044; initially C. fungifaber; NHMUK 13671870 • soldier; Mbalmayo; 3°27′ N, 11°29′ E; Mar. 1996; P. Eggleton et al. leg.; study code: DJ U046; initially C. fungifaber; NHMUK 13671859 • soldier; Mbalmayo; 3°27′ N, 11°29′ E; Aug. 1996; P. Eggleton et al. leg.; study code: DJ U043; initially C. fungifaber; NHMUK 13671873 • soldier; Nsimi; 3°10′ N, 11°58′ E; Nov. 1996; M. Harry leg.; study code: DJ P136; GenBank no PQ679252 (mitogenome); MNHN EP9869 • soldier; Nsimi; 3°10′ N, 11°58′ E; Nov. 1996; M. Harry leg.; study code: DJ P137; MNHN EP9870 • soldier; Nsimi; 3°10′ N, 11°58′ E; Nov. 1996; M. Harry leg.; study code: DJ P138; MNHN EP9871 • soldier; Nsimi; 3°10′ N, 11°58′ E; Nov. 1996; M. Harry leg.; study code: DJ P139; MNHN EP9872 • soldier; Nsimi; 3°10′ N, 11°58′ E; Nov. 1996; M. Harry leg.; study code: DJ P140; MNHN EP9873 • soldier, worker, ♂ (alate); Makak; 3°38′ N, 11°4′ E; 27 Nov. 2013; J. Birkett-Smith and J. Dahl leg.; study code: DJ 0650; initially C. fungifaber; NHMUK 13671853 • soldier, worker, ♂ (alate); Bipindi; 3°5′ N, 10°24.5′ E; 23 Nov. 2016; P. Akama leg.; study code: DJ 0452; GenBank nos MN646713 (COI), MN685915 (COII), MN685976 (28S), PQ679181 (mitogenome); BE RMCA INS.Iso.059200 • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 23 Nov. 2016; P. Akama leg.; study code: DJ 0468; GenBank no PQ679188 (mitogenome); BE RMCA INS.Iso.059196 • soldier, worker; Bipindi; 3°5′ N, 10°24.5′ E; 23 Nov. 2016; P. Akama leg.; study code: DJ 0469; GenBank no PQ679195 (mitogenome); BE RMCA INS. Iso.059195 • soldier, worker, ♀ (alate); Mbongé; 4°32.2′ N, 9°6.7′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0450; GenBank nos MN646712 (COI), MN685914 (COII), MN685975 (28S); BE RMCA INS. Iso.059194 • soldier, worker; Mbongé; 4°32.2′ N, 9°6.7′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0451; GenBank no PQ679193 (mitogenome); BE RMCA INS.Iso.059202 • soldier, worker, ♂ (alate); Mbongé; 4°32.2′ N, 9°6.7′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0466; GenBank no PQ679246 (mitogenome); BE RMCA INS.Iso.059198 • soldier, worker, ♂ (alate); Mbongé; 4°32.2′ N, 9°6.7′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0467; GenBank no PQ679226 (mitogenome); BE RMCA INS. Iso.059203 • soldier, worker, ♀ (queen); Ebogo; 3°22.894′ N, 11°27.862′ E; 23 May 2017; Y. Roisin leg.; study code: DJ 0506; GenBank nos MN646714 (COI), MN685916 (COII), MN685977 (28S); BE RMCA INS.Iso.059206 • soldier, worker; Ebogo; 3°22.872′ N, 11°27.796′ E; 6 Jun. 2017; Y. Roisin leg.; study code: DJ 0521; GenBank no PQ679222 (mitogenome); BE RMCA INS.Iso.059205 • soldier, worker; Mbalmayo; 3°28.841′ N, 11°34.646′ E; 6 Jun. 2017; Y. Roisin leg.; study code: DJ 0520; GenBank no PQ679185 (mitogenome); BE RMCA INS.Iso.059207

CÔTE D’IVOIRE • soldier; Banco N.P.; 5°23′ N, 4°3′ W; 1 Jan. 1934; P.-P. Grassé leg.; study code: DJ P199; MNHN EP9849 • soldier; Banco N.P.; 5°23′ N, 4°3′ W; 8 Jan. 1934; P.-P. Grassé leg.; study code: DJ P153; MNHN EP9844 • soldier, worker; Banco N.P.; 5°24′ N, 4°3′ W; 9 Jan. 1934; P.-P. Grassé leg.; study code: DJ 0203; initially C. fungifaber in MNHN; MNHN EP9840 • soldier; Banco N.P.; 5°23′ N, 4°3′ W; Jan. 1939; P.-P. Grassé leg.; study code: DJ P157; MNHN EP9845 • soldier, ♀ (queen); Banco N.P.; 5°23′ N, 4°3′ W; 6 Feb. 1947; C. Noirot leg.; study code: DJ P194; MNHN EP9848 • soldier, ♂ (king); Adiopodoumé; 5°19.75′ N, 4°8′ W; 22 May 1947; C. Noirot leg.; study code: DJ P193; MNHN EP9847 • soldier; Banco N.P.; 5°19′ N, 4°10′ W; 26 Jul. 1947; C. Noirot leg.; study code: DJ P325; MNHN EP9863 • soldier; Mambo; 5°46′ N, 4°3′ W; 19 Oct. 1947; C. Noirot leg.; study code: DJ P324; MNHN EP9862 • soldier; Mambo; 5°46′ N, 4°3′ W; 19 Oct. 1947; C. Noirot leg.; study code: DJ P333; MNHN EP9948 • soldier; Mambo; 5°46′ N, 4°3′ W; 20 Oct. 1947; C. Noirot leg.; study code: DJ P334; MNHN EP9949 • soldier; Yapo; 5°44′ N, 4°5′ W; 10 Oct. 1947; C. Noirot leg.; study code: DJ P326; MNHN EP9864 • soldier, worker, ♀ (alate); Petit Yapo réserve forestière; 5°46′ N, 4°8′ W; 22 Aug. 1953; M. Lüscher leg.; study code: DJ 0277; initially C. fungifaber; AMNH • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 6 Dec. 1959; C. Noirot leg.; study code: DJ P211; MNHN EP9860 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 25 Dec. 1959; C. Noirot leg.; study code: DJ P208; MNHN EP9857 • soldier, worker, ♂ (alate); near Abidjan; 5°23′ N, 4°3′ W; ca 1960; C. Noirot leg.; study code: DJ 0268; initially C. fungifaber; AMNH • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 17 Jan. 1960; C. Noirot (?) leg.; study code: DJ P203; MNHN EP9852 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 17 Jan. 1960; C. Noirot (?) leg.; study code: DJ P205; MNHN EP9854 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 17 Jan. 1960; C. Noirot (?) leg.; study code: DJ P207; MNHN EP9856 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 17 Jan. 1960; C. Noirot leg.; study code: DJ P209; MNHN EP9858 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 17 Jan. 1960; C. Noirot leg.; study code: DJ P210; MNHN EP9859 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 30 Jan. 1960; C. Noirot (?) leg.; study code: DJ P201; MNHN EP9850 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 30 Jan. 1960; C. Noirot (?) leg.; study code: DJ P204; MNHN EP9853 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 20 Mar. 1960; C. Noirot leg.; study code: DJ P329; MNHN EP9866 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 7 May 1960; C. Noirot (?) leg.; study code: DJ P202; MNHN EP9851 • soldier, ♂ (alate); Banco N.P.; 5°24′ N, 4°3′ W; 7 May 1960; C. Noirot (?) leg.; study code: DJ P206; MNHN EP9855 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 12 Jun. 1960; C. Noirot leg.; study code: DJ P330; MNHN EP9867 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 17 Jun. 1960; C. Noirot leg.; study code: DJ P336; MNHN EP9951 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 9 Oct. 1960; C. Noirot leg.; study code: DJ P327; MNHN EP9865 • soldier; Banco N.P.; 5°24′ N, 4°3′ W; 9 Oct. 1960; C. Noirot leg.; study code: DJ P331; MNHN EP9868 • soldier, ♂ (king); Abidjan; 5°20′ N, 3°59′ W; 29 Dec. 1960; C. Noirot leg.; study code: DJ P347; MNHN • soldier, ♂ (alate); Adiaké; 5°22′ N, 3°19.7′ W; 15 Apr. 1963; C. Noirot leg.; study code: DJ P348; MNHN • soldier; Banco N.P.; 5°23′ N, 4°3′ W; 21 Aug. 1968; G. Josens leg.; study code: DJ U001; initially C. modestior; NHMUK 13671927 • soldier, worker; Banco N.P.; 5°24′ N, 4°3′ W; 21 Aug. 1968; G. Josens leg.; study code: DJ 0202; initially C. modestior in RMCA; BE RMCA INS.Iso.059204 • soldier, worker; Ahoutoué; 5°28′ N, 3°45′ W; 9 Aug. 1969; Vincent leg.; study code: DJ 0229; BE RMCA INS.Iso.059209 • soldier, worker; Banco N.P.; 5°23.005′ N, 4°3.341′ W; 17 Feb. 2015; G. Josens leg.; study code: DJ 0412; GenBank nos MN646711 (COI), MN685913 (COII), MN685974 (28S), PQ679172 (mitogenome); BE RMCA INS.Iso.059208 .

EQUATORIAL GUINEA • soldier; Punta Frailes; 3°47′ N, 8°43′ E; ca 1894; R.M. Downes leg.; study code: DJ U110; initially C. gaigei; NHMUK 13671887 • soldier, worker, ♀ (alate); Punta Frailes; 3°47′ N, 8°43′ E; Oct. 1901; L. Fea leg.; study code: DJ 0311; initially C. fungifaber; MCGD .

GABON • soldier; Mekambo; 1°1′ N, 13°56′ E; Jan. 1957; P.-P. Grassé and C. Noirot leg.; study code: DJ P167; MNHN EP9846 • soldier, worker; Piste du Bouéni; 0°13′ N, 11°48′ E; 1 Jan. 1957; P.-P. Grassé and C. Noirot leg.; study code: DJ 0855; MNHN EP9843 • soldier, worker; Piste du Bouéni, km 53; 0°44′ N, 13°12′ E; 1 Jan. 1957; P.-P. Grassé and C. Noirot leg.; study code: DJ 0853; MNHN EP9841 • soldier, worker, ♂ (king), ♀ (queen); forêt dense de Djidji; 0°13′ N, 11°48′ E; 25 Jan. 1957; P.-P. Grassé and C. Noirot leg.; study code: DJ 0854; MNHN EP9842 • soldier, worker; La Lopé N.P.; 0°15′ S, 11°35′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ P242; MNHN EP9875 • soldier, worker; La Lopé N.P.; 0°12.7′ S, 11°33.5′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0623; GenBank nos DQ127302 (COII), DQ246527 (ITS2), PQ679189 (mitogenome); initially C. aff. subarquatus “spA” (cf. Roy et al. 2006); MNHN EP9878 • soldier, worker; Doda (north of La Lopé N.P.); 0°4.5′ S, 11°25.5′ E; 10 Mar. 1998; M. Harry leg.; study code: DJ 0625; GenBank nos DQ246541 (COII), DQ246526 (ITS2) initially C. aff. subarquatus “spD” (cf. Roy et al. 2006); MNHN EP9879 • soldier, ♀ (queen); La Lopé N.P.; 0°15′ S, 11°35′ E; 13 Mar. 1998; M. Harry leg.; study code: DJ P243; MNHN EP9876 • soldier, worker; La Lopé N.P.; 0°12.7′ S, 11°33.5′ E; 13 Mar. 1998; M. Harry leg.; study code: DJ 0626; GenBank nos DQ127299 (COII), DQ246529 (ITS2), PQ679214 (mitogenome); initially C. aff. subarquatus “spA” (cf. Roy et al. 2006); MNHN EP9880 • soldier; La Lopé N.P.; 0°9′ S, 11°36′ E; 6 Mar. 1998; M. Harry leg.; study code: DJ P232; MNHN EP9874 • soldier, worker; Bitam; 2°5′ N, 11°30′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0456; BE RMCA INS.Iso.059197 • soldier, worker; Bitam; 2°5′ N, 11°30′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0457; BE RMCA INS.Iso.059201 • soldier; Bitam; 2°5′ N, 11°30′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0458; BE RMCA INS.Iso.059199 • soldier, worker; Nkobissimo [= Nkolmengboua?]; 2°14.3′ N, 11°29.25′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0453; GenBank no PQ679216 (mitogenome); BE RMCA INS. Iso.059193 • soldier, worker; Ipassa; 0°30.66′ N, 12°48′ E; 27 Jan. 2025; J. Šobotnik leg.; study code: DJ 0974; BE RMCA INS.Iso.059949 .

GHANA • soldier, ♀ (queen); Bobiri Forest Reserve; 6°40′ N, 1°21′ W; 21 Feb. 1959; W.A. Sands leg.; study code: DJ U114; initially C. gaigei; NHMUK 13671881 .

NIGERIA • soldier; Ijebu–Ode road; 6°50′ N, 3°54′ E; 2 May 1950; G.C. Webb leg.; study code: DJ N126; initially C. fungifaber; AMNH • soldier; North of Lagos; 6°49′ N, 3°13′ E; 11 Feb. 1955; W.V. Harris leg.; study code: DJ U063; initially C. fungifaber; NHMUK 13671865 • soldier; Mofu River forest reserve; 6°10′ N, 7°10′ E; 15 Feb. 1955; W.V. Harris leg.; study code: DJ U062; initially C. fungifaber; NHMUK 13671864 • soldier; Port Harcourt; 4°46′ N, 7°1′ E; 5 Dec. 1955; D. Kay leg.; study code: DJ U057; initially C. fungifaber; NHMUK 13671868 • soldier; Sapele; 5°53′ N, 5°41′ E; 31 Jan. 1956; D. Kay leg.; study code: DJ U056; initially C. fungifaber; NHMUK 13671866 • soldier; Obanikoro; 6°33′ N, 3°22′ E; 8 Jan. 1957; W. Wilkinson leg.; study code: DJ U034; initially C. fungifaber; NHMUK 13671861 • soldier; Ugo; 6°5′ N, 6°0′ E; 11 Jan. 1957; W. Wilkinson leg.; study code: DJ U061; initially C. fungifaber; NHMUK 13671860 • soldier, worker; Ilaro (near)?; 7°26′ N, 3°5′ E; 6 Apr. 1957; W. Wilkinson leg.; study code: DJ 0549; initially C. fungifaber; NHMUK • soldier, ♀ (queen); Calabar-Mamfe road; 5°17′ N, 8°34′ E; 22 Apr. 1957; W. Wilkinson leg.; study code: DJ U035; initially C. fungifaber; NHMUK 13671863 • soldier; Port Harcourt–Owerri road; 5°11′ N, 6°52′ E; 19 Jun. 1957; W. Wilkinson leg.; study code: DJ U064; initially C. fungifaber; NHMUK 13671867 • soldier, worker; Port Harcourt–Owerri road; 5°9.5′ N, 6°51′ E; 19 Jun. 1957; W. Wilkinson leg.; study code: DJ 0548; initially C. fungifaber; NHMUK • soldier; Mamu forest reserve; 6°11′ N, 7°10′ E; 28 Feb. 1958; W.A. Sands leg.; study code: DJ U058; initially C. fungifaber; NHMUK 13671862 • soldier; Enugu – Otukpo road; 6°45′ N, 7°28′ E; 5 Mar. 1958; W.A. Sands leg.; study code: DJ U055; initially C. fungifaber; NHMUK 13671858 • soldier; Ayangba; 7°30′ N, 7°10′ E; 6 Mar. 1958; W.A. Sands leg.; study code: DJ U059; initially C. fungifaber; NHMUK 13671857 • soldier; Ijebu–Ode road; 6°49′ N, 3°55′ E; 12 Dec. 1959; W.A. Sands leg.; study code: DJ U060; initially C. fungifaber; NHMUK 13671859 .

Supplementary material of dubious I. fungifaber

Four samples from the CAR, near Bangui, and two from eastern DRC (Fig. 45) show a soldier’s morphology entirely consistent with that of I. fungifaber . As these samples come from semi-deciduous forest and forest galleries (which is not characteristic of I. fungifaber) and as it was not possible to sequence them, these six samples, are presented here as “dubious, possibly cryptic I. fungifaber ”. The distance between Bangui and the nearest harvesting point in Gabon is around 500 km, and between Bangui and eastern DRC, the distance is around 1300 km.

Of the six old museum samples examined, four samples were initially labelled as C. fungifaber and two not identified.

CENTRAL AFRICAN REPUBLIC • soldier; route Mbaïki–Bagandou; 3°52′ N, 17°52′ E; 21 Jun. 1948; P.-P. Grassé and C. Noirot leg.; study code: DJ P188; MNHN EP9825 • soldier; Boukoko; 3°54′ N, 17°55′ E; 2 Jul. 1948; P.-P. Grassé and C. Noirot leg.; study code: DJ P189; MNHN EP9826 • soldier; Bangui; 4°27′ N, 18°32′ E; 1975; Becker leg.; study code: DJ U037; initially C. fungifaber; NHMUK 13671856 • soldier; Bangui; 4°27′ N, 18°32′ E; 1975; Becker leg.; study code: DJ U038; initially C. fungifaber; NHMUK 13671855 .

DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (queen); “ Alto Uele, foresta”; 2°30′ N, 29°30′ E; [no date]; S. Patrizi leg.; study code: DJ 0312; initially C. fungifaber in MCGD • soldier, ♀ (alate); Epulu; 1°24′ N, 28°34′ E; 15 May 1948; A. Emerson leg.; study code: DJ N129; initially C. fungifaber in AMNH .

Historical review

The alate imago from Bonge [= Mbongé], Cameroon, was briefly described by Sjöstedt (1896: 297) under the name Eutermes fungifaber, mentioning it as very close to Eutermes mordax (Smeathman, 1781) .

Four years later, Sjöstedt (1900: 143–149) provided more detailed descriptions (text and figures) of all castes and nest. Sjöstedt (1926: 218–225) included Cubitermes fungifaber in imagines' and soldiers' keys of Cubitermes species and inserted it in a “ fungifaber -Gruppe” of species with a not forward humped soldier’s frons.

Emerson (1928: 517–519) referred to this species as Mirotermes (Cubitermes) fungifaber; he provided further descriptions and measurements of all castes, based on samples collected by G. Zenker in Cameroon.

Snyder (1949: 159) catalogued this species under the name Cubitermes fungifaber in the sub-family Termitinae .

Bouillon & Vincke (1971: 269) described the enteric valve of C. fungifaber as belonging to the first of three types, the “simple type ” without any spatula.

Cubitermes fungifaber was considered as a senior synonym of (a) C. banksi, (b) C. comstocki and (c) C. schmidti (Emerson unpublished “Card catalog”; Ruelle 1992: 501; Krishna et al. 2013: 1913, 1918, 1935). These three species are somewhat smaller than C. fungifaber, but synonymies are compatible with their enteric valve patterns (Josens & Deligne 2019: 39–42).

To obtain genetic sequences of I. fungifaber, one of us (PA) went and collected samples at Mbongé, type location of I. fungifaber, and at Bipindi, type location of I. banksi, I. comstocki and I. schmidti; some of the soldiers in those samples were as small as the soldiers of C. comstocki and Hellemans et al. (2021: 233) confirmed their synonymies and placed the species in the restored genus Isognathotermes .

Cubitermes fungifaber was considered a senior synonym of C. fungifaber var. elongata (Krishna et al. 2013: 1923) and a possible senior synonym of C. planifrons (Emerson in his unpublished “card catalog”, quoted by Krishna et al. 2013: 1931) but these synonymies are rejected because the two taxa proposed as junior synonyms do not have enteric valves of the fungifaber pattern, but well of the finitimus one (Josens & Deligne 2019: 42–44).

Cubitermes aff. subarquatus “spA” and Cubitermes aff. subarquatus “spD” were mentioned by Roy et al. (2006) in La Lopé National Parc, Gabon; on a genetical base they were considered as cryptic species. Hellemans et al. (2021: 231) placed these species in the restored genus Isognathotermes as synonyms of I. fungifaber .

As already mentioned, the geographic distribution of I. fungifaber is clearly dependant on humid, forest environments. In at least one documented case, its geographic distribution has been changing during the second half of the 20 th century, possibly in relation with global and local climate change. Isognathotermes fungifaber used to be common in the Banco National Park near Abidjan, RCI: see the publications by Noirot et al. (1986), Han & Lepage (1991), and personal observation (GJ, in 1970); however, nowadays, this national park is almost totally included within the city of Abidjan, which presumably influences the local climate. Moreover, the annual rainfall in Côte d’Ivoire has been declining by about 20% during the last decades, mainly during the late sixties and early seventies (Servat et al. 1999), possibly linked with huge deforestation. This might explain that two of us (GJ and YR) could hardly find a single nest of this species in 2015 in the Banco National Park.

Redescription

Imago

COLOUR. Head capsule: well sclerotised, fresh samples very dark, C7–C8 (C6–C 7 in long preserved imagines); fontanelle concolorous or almost so with head capsule. Postclypeus C6–C7 slightly paler (one level) than head capsule. Antennae C5–C6 without any difference between proximal and distal articles. Thorax: pronotum C6–C8 (C5–C 7 in long preserved imagines), as postclypeus, generally one level paler than head capsule; meso- and metanotum C5–C7, sometimes somewhat paler than pronotum. Legs C3–C5; tibia usually slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf3–Cf4), anterior veins darker. Abdomen: tergites C5–C7. Sternites appreciably paler in middle (C3– C4) with both sides darker (C4–C6); posterior sternites darker (C4–C6) than anterior.

SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 ×, sometimes only at 80 ×. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 5–12 on the carina of fore coxa and 1–8 on the ventral side of fore coxa and trochanter; tibia pilose fore, mid, and hind tibia furnished with 30–40 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards.

STRUCTURE (measurements in Table 5; Fig. 42). Size: the imagines of I. fungifaber are, on average, among the smallest of the genus Isognathotermes (Fig. 26). Head capsule: compound eyes shortly oval; ocelli shortly oval to oval, removed from eyes by a distance equal to 0.7–1.4 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: generally 16 articles on alate individuals, rarely 17 (one individual out of 16), always shortened by amputation by three to four articles in queens and kings. Labrum: cupola shaped, wider than long. Left mandible with apical tooth on average longer than in most other species of the genus (Fig. 27) and always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a more or less sharp carina. Wings: R1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 3–5 and 7–11 branches respectively. Gut (only studied in some cases): enteric valve weakly developed and hardly comparable with workers’ and soldiers’ valves; odd PCs are slightly longer than even PCs (Fig. 6). Caecum present as a small amorphous button.

* Abbreviations: see definitions in Material and methods.

Soldier

COLOUR. Head capsule generally tending towards deep palette (Cd4–Cd6) becoming fader and darker (e.g., Cf5–Cf7) in long preserved samples; there is always a gradient from a darker frons to a paler back (e.g., from C6–C4), smooth and extended in 55% of the samples examined but more abrupt in the remaining 45% giving in some cases the impression that the head capsule is bicolorous (as in Fig. 13). Antennae and labrum sometimes concolorous or one level paler than head capsule. Mandibles dark (C6–C8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons; in long preserved samples, this clearing tends to disappear. Thorax and legs paler than head capsule (C2–C4) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellow to reddish-brown tinge on tergites.

SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with some large setae on lobes. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta but not any spines (in 68% of the samples examined) or furnished with 1–3 spines on carina and 0–2 on ventral side; trochanter generally with some long lined-up setae, sometimes with six or seven spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6–15 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards.

STRUCTURE (measurements in Table 5; Fig. 43). Size: the soldiers of I. fungifaber are among the smallest of the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with (in 86% of the samples examined) a narrowing near posterior third or fourth; from antennal sockets sides converge more or less clearly towards bases of mandibles; posterior side regularly rounded or sometimes (in 29% of the samples examined) with a short straight or even concave part in the middle. In profile: upper profile almost always slightly concave; angle between extended mandibles and frons varies from right to a little obtuse; frons with a sketched or without any anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part either roundly convex or forming an acute widening or even a kind of ear on each side. Antennae: of 14.5–15 articles. Labrum: always deeply bifurcate and wider than long, with sides varying from lyre-shaped (in 84% of the samples examined) to slightly convex; lobes angular, frequently with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like strongly curved (on average, I. fungifaber, and I. planifrons, have the highest left mandible apical curvature index in the genus Isognathotermes); inner edges generally smooth with one distinct but small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, as wide as 55–70% of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather small, best seen in ventral view, near centre of abdomen, lobed, sometimes (in 18% of the samples examined) with a lobe expanded somewhat forward. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 20% longer than the even cushions, generally without any crest or with crests very weakly developed.

Worker

COLOUR. Head capsule pale (C1–C3) turning grey in long preserved samples. Antennae: proximal articles pale (C2–C3), distal articles always one to two levels darker (C4–C5). Thorax, nota, and legs pale (C1–C3). Abdomen grey to red-brown owing to digestive bolus.

SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3–5 spines on carina and 1–4 on ventral side; fore trochanter with 5–7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards.

STRUCTURE (measurements in Table 5, Fig. 44). Size: the workers of I. fungifaber are, on average, among the smallest of the genus Isognathotermes (but with large overlapping on several other species: Fig. 31). Head capsule: weakly sclerotised (except mandibles). Antennae of 14.5 (rarely 14) articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal tooth faint (visible as an undulated edge and disappearing in worn mandibles), third marginal tooth with a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 57–70% of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the fungifaber pattern with triradial symmetry: the odd PCs, in their downstream part, are rather narrow and bear crests that are as high as or higher than they are wide, with long and strong bristles; supporting bristles are generally numerous: 14–38 on each side of the odd PCs; only three samples (out of 54) show an intermediate EVA between the fungifaber and finitimus patterns; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, generally with two or three (up to five) short lobes, sometimes (in 14% of the samples examined) with a lobe expanded forward.

Chorology-ecology

The geographic distribution of I. fungifaber is clearly linked with humid, evergreen forests generally at less than 300 km from the Atlantic coast (Fig. 45): it includes from west to east the eastern Guinean forests, the Cross-Sanaga-Bioko coastal forests, the Atlantic Equatorial coastal forests, and the Northwestern Congolian lowland forests ecoregions. This distribution partly resembles that of I. planifrons (that has a different EVA).

However, some samples of “dubious I. fungifaber ” come from the northeastern Congolian lowland forests ecoregion without it being possible to guarantee the true species of these samples.

Molecular data

Thirteen mitogenomes of I. fungifaber are published alongside this work (GenBank accessions: see Supp. file 3 and ‘Material examined’). These were collected from RCI, Cameroon, and Gabon. Most samples exhibit less than 1% dissimilarity with each other (Supp. file 4); except for sample DJ 0451 being the most dissimilar (1.50%) collected in one of the type localities of the species, and for sample DJ 0412, collected at the westernmost part of its distribution (1.32%). Our phylogenetic reconstructions based on the COII gene (Figure SF2) includes samples identified as “ Cubitermes aff. subarquatus spA” (DJ 0623 and DJ 0626) and “spD” (DQ246541, DJ 0625) of Roy et al. (2006). The two mitogenomes of “spA” samples were successfully re-sequenced from the samples having produced the COII sequences published under accession DQ127302 and DQ127299 but not that of the “spD” sample. Therefore, the cryptic species status of the “spD” proposed by Roy et al. (2006) cannot be discussed here.

Remarks

Isognathotermes fungifaber is quite variable in size and much more in soldiers and workers than in imagines: we can deduct from Table 5 that variability in head width for I. fungifaber is 14% in imagines, but 31% in soldiers and 28% in workers

The soldiers of several samples have an intermediate morphology between I. fungifaber and I. severus (having the same kind of EVA): the easiest way to distinguish them is their origin since they seem to be mutually exclusive. This is linked with their preferred habitat: on the one hand, I. fungifaber seems to be restricted to evergreen forests with, however, one known exception: DJ 0625, that was proposed as a cryptic species (“spD”) by Roy et al. 2006: it was collected in the Doda gallery forest, surrounded with savannahs, north of La Lopé National Park, Gabon. On the other hand, I. severus is common in savanna landscapes of West and Central Africa (from Gambia to CAR) but also in orchards, secondary forests and, in some regions, it is also found in evergreen forest but only in places where I. fungifaber is absent as for example in RCI near the border with Liberia.